Grain yield of wheat (Triticum aestivum L.) under long‐term heat stress is sink‐limited with stronger inhibition of kernel setting than grain filling

The aim of the study was to investigate source‐sink relations of wheat under continuous heat stress and to identify bottle necks of yield formation. A pot experiment was conducted in two climatic chambers exposing wheat plants (Triticum aestivum L. cv. Thasos) either to day/night temperatures of 20/20°C (control conditions) or of 30/25°C (heat stress) during the whole vegetation period in the absence of plant water deficit. Plants were harvested at four phenological stages: three‐node stage (DC 33), start of flowering (DC 61), grain filling (DC 75) and maturity (DC 94). Heat stress shortened the development phases of the plants and caused a significant decrease in total above‐ground biomass between 19% and 41%. At grain filling and at maturity, the reductions in total shoot biomass mainly resulted from grain yield depressions by 77% and 58%, respectively. The ear number per plant was significantly higher under heat stress in comparison with the control, at maturity it was more than doubled. On the contrary, under heat stress, the kernel number per ear was strongly decreased by 83% and 75% during grain filling and at maturity, respectively. The decrease in individual kernel weight was 23% at maturity. Thus, the heat‐stressed plants were able to strongly increase the number of ear‐bearing tillers which were able to set only a small number of kernels, yet these kernels showed good grain filling. The harvest index (HI) of heat‐stressed plants was significantly reduced by 36% (control: HI = 50.1% ± 0.4, heat: HI = 32.2% ± 0.9***). The plants in the stress treatment adapted to the adverse conditions by less biomass production which presumably allowed a higher transpiration without an increase in total water consumption. Nevertheless, under heat stress, the water use efficiency (WUE_grain) was strongly decreased by 62% as a result of a small grain yield. In ears and grains, the sucrose, glucose and fructose concentrations were not significantly different between control and heat stress at start of flowering and during grain filling. Thus, the supply of assimilates was not restricted (no source limitation). Sink capacity was reduced by heat stress, as lesser and smaller kernels were produced than in the control. Concerning sink activity, the sink‐limiting step during kernel set is probably the active transport of hexoses across the plasma membrane into the developing kernels, which could also affect grain filling. This needs to be investigated in more detail in further studies.     

Bread and Durum Wheat under Heat Stress: A Comparative Study on the Photosynthetic Performance

The photosynthetic responses to heat stress, during grain filling, in four genotypes of Triticum aestivum L. (Sever and Golia) and Triticum turgidum subsp. durum (Acalou and TE 9306), chosen according to its genetic background diversity, were investigated. All wheat genotypes (excepting Golia) showed synergistic trends implicating the internal CO₂ concentration, net photosynthesis and stomatal conductance. Additionally, the modifications of net photosynthesis were associated with changes in stomatal control. Chlorophyll a fluorescence parameters (minimal fluorescence, maximal and variable fluorescence, intrinsic efficiency of PSII in darkness, non‐photochemical quenching, photochemical quenching and energy‐dependent chlorophyll fluorescence quenching) further pointed heat protective mechanisms, implicating F_v/F_m stabilization (i.e. maintaining the efficiency of PS II) and electron transport rate preservation. It is concluded that, comparatively to bread wheat, the photosynthetic performance of durum wheat is more tolerant to heat stress, as stomatal conductance and transpiration are less affected.     

The Alleviating Effect of Elevated CO2 on Heat Stress Susceptibility of Two Wheat (Triticum aestivum L.) Cultivars

This study analysed the alleviating effect of elevated CO₂ on stress‐induced decreases in photosynthesis and changes in carbohydrate metabolism in two wheat cultivars (Triticum aestivum L.) of different origin. The plants were grown in ambient (400 μl l^?1) and elevated (800 μl l^?1) CO₂ with a day/night temperature of 15/10 °C. At the growth stages of tillering, booting and anthesis, the plants were subjected to heat stress of 40 °C for three continuous days. Photosynthetic parameters, maximum quantum efficiency of photosystem II (PSII) photochemistry (F_v/F_m) and contents of pigments and carbohydrates in leaves were analysed before and during the stress treatments as well as after 1 day of recovery. Heat stress reduced P_N and F_v/F_m in both wheat cultivars, but plants grown in elevated CO₂ maintained higher P_N and F_v/F_m in comparison with plants grown in ambient CO₂. Heat stress reduced leaf chlorophyll contents and increased leaf sucrose contents in both cultivars grown at ambient and elevated CO₂. The content of hexoses in the leaves increased mainly in the tolerant cultivar in response to the combination of elevated CO₂ and heat stress. The results show that heat stress tolerance in wheat is related to cultivar origin, the phenological stage of the plants and can be alleviated by elevated CO₂. This confirms the complex interrelation between environmental factors and genotypic traits that influence crop performance under various climatic stresses.     

Traits in Spring Wheat Cultivars Associated with Yield Loss Caused by a Heat Stress Episode after Anthesis

Heat stress resulting from climate change and more frequent weather extremes is expected to negatively affect wheat yield. We evaluated the response of different spring wheat cultivars to a post‐anthesis high temperature episode and studied the relationship between different traits associated with heat tolerance. Fifteen spring wheat (Triticum aestivum L.) cultivars were grown in pots under semifield conditions, and heat stress (35/26 °C) and control treatments (20/12 °C) were applied in growth chambers for 5 days starting 14 days after flowering. The heat stress treatment reduced final yield in all cultivars. Significant variation was observed among cultivars in the reduction in average grain weight and grain dry matter yield under heat stress (up to 36 % and 45 %, respectively). The duration of the grain‐filling period was reduced by 3–12 days by the heat treatment. The reduction in the grain‐filling period was negatively correlated with grain nitrogen yield (r = ?0.60). A positive correlation (r = 0.73) was found between the treatment effect on green leaf area (GLA) and the reduction in yield resulting from heat stress. The amount of stem water‐soluble carbohydrates (WSC) was not related to treatment effects on grain yield or grain weight. However, the treatment effect on stem WSC remobilization was negatively correlated with reduction in grain‐filling duration due to heat stress (r = ?0.74) and positively with treatment effect on grain N yield (r = 0.52). The results suggest that the effect of the heat treatment on GLA was the trait most associated with yield reduction in all cultivars. These findings suggest the importance of ‘stay green’‐associated traits in plant breeding as well as the need for better modelling of GLA in crop models, especially with respect to brief heat episodes during grain filling. There is in particular a need to model how heat and other stresses, including interacting effects of heat and drought, affect duration of GLA after flowering and how this affects source–sink relations during grain filling.     

Independent and Combined Effects of High Temperature and Drought Stress During Grain Filling on Plant Yield and Chloroplast EF‐Tu Expression in Spring Wheat

High temperature and drought stress are among the two most important environmental factors influencing crop growth, development and yield processes. These two stresses commonly occur in combination. Objectives of this research were to investigate the independent and combined effects of high temperature and drought stress during grain filling on physiological, vegetative and yield traits and expression of a chloroplast protein synthesis elongation factor (EF‐Tu) of wheat (Triticum aestivum L.). Two spring wheat cultivars (Pavon‐76 and Seri‐82) were grown at control temperatures (CT; day/night, 24/14 °C; 16/8 h photo/dark period) from sowing to heading. Thereafter, one half of the plants were exposed to high temperature stress (HT; 31/18 °C in Exp. 1 and 34/22 °C in Exp. 2), drought stress (withholding water), or a combination of both HT and drought stress. There were significant influences of HT and/or drought stress on physiological, growth and yield traits. There was no cultivar or cultivar by temperature or cultivar by drought interaction effects on most traits. The decreases in leaf photosynthesis were greater at HT compared with drought alone throughout the stress period, and the combination of HT and drought had the lowest leaf photosynthetic rates. Overall, HT or drought had similar effects (about 48–56 % decrease) on spikelet fertility, grain numbers and grain yield. High temperature decreased grain numbers (by 56 % averaged across both experiments) and individual grain weight (by 25 %), while, respective decreases due to drought were 48 % and 35 %. This suggests that the grain numbers were more sensitive to HT and grain weights to drought for the range of temperatures tested in this research. The interaction between HT and drought stress was significant for total dry weights, harvest index and spikelet fertility, particularly when HT stress was severe (34/22 °C). The combined effects of HT and drought were greater than additive effects of HT or drought alone for leaf chlorophyll content, grain numbers and harvest index. High temperature stress and the combination of HT and drought stress but not drought stress alone resulted in the overexpression of EF‐Tu in both spring wheat cultivars.     

Glycinebetaine Improved Photosynthesis in Canola under Salt Stress: Evaluation of Chlorophyll Fluorescence Parameters as Potential Indicators

Salinity is a widespread soil problem limiting productivity of oilseed crops worldwide. Reduction in growth generally associated with decline in photosynthesis. Accumulation of osmo‐protectants such as glycinebetaine (GB) permits the plant to cope with salt stress. This study was aimed to understand the mechanism by which exogenous GB application improves photosynthetic capacity and salt tolerance in two diverse lines of canola differing in salt tolerance using chlorophyll fluorescence technique. Glycinebetaine was applied as foliarly or through rooting medium to two canola lines Dunkeld and Cyclone that were grown under non‐saline or saline conditions. Root‐applied GB caused adverse effects whereas foliar application of GB was effective in improving growth of canola cultivars. However, the response of salt‐tolerant cultivar Dunkeld was stronger than that of salt sensitive Cyclone. Foliar application of GB improved accumulation of proline and plant water status. Application of GB enhanced the photosynthetic CO₂ fixation, stomatal conductance (g_s) and water‐use efficiency. The tolerant lines Dunkeld had more responsive to GB application. Chlorophyll fluorescence measurements (F_o, F_m, F_v, F_o/F_m, F_v/F_o and F_v/F_m ratios) revealed that salt stress reduced energy trapping efficiency by damaging oxygen evolving complex, over reduction of Q_A resulting in occurrence of chronic photoinhibition. However, exogenous GB protected the oxygen evolving centre of PSII and maintains activity of PSII. Although root‐applied GB adversely affected the growth of canola plants, it did not have any adverse effect on PSII photochemistry. Moreover, fluorescence parameters could provide a rapid means for determining salt tolerance in canola and could be a rapid and sensitive test to identify genotypes highly tolerant to salt stress.     

Differential root responses in two cultivars of winter wheat (Triticum aestivum L.) to elevated ozone concentration under fully open‐air field conditions

The effect of elevated tropospheric ozone concentration [O₃] on root processes in wheat systems of different O₃ sensitivity is not well understood. Two wheat cultivars (cv. Y15 and YN19) with contrasting O₃ tolerance were grown in a fully open‐air O₃ enrichment platform for one season. We found that elevated O₃ (EO₃) (50% above the ambient O₃) significantly decreased the total biomass at all key growth stages and the yield of the O₃‐sensitive cultivar YN19 but did not affect those of the O₃‐tolerant cultivar Y15. EO₃ significantly decreased the root biomass of two wheat cultivars at the jointing and grain‐filling stages. EO₃ significantly decreased the root length, length density, surface area and volume of the two cultivars at the jointing stage but increased those of YN19 at the grain‐filling stage. EO₃ significantly increased the root activities (specific root respiration rates) of Y15 and YN19 at the jointing, heading and grain‐filling stages. EO₃ significantly decreased the contribution of fresh root respiration to soil respiration (CRS) of YN19 at the jointing stage but increased it at the heading stage; however, it did not change the CRS of Y15 at any growth stages. This study indicates that the effects of EO₃ on root morphology and activity varied among wheat cultivars, and suggest that we can breed O₃‐tolerant cultivars to maintain crop yield under higher [O₃] scenarios.     

Does post‐anthesis heat stress affect plant phenology,physiology, grain yield and protein content of durum wheat in a semi‐arid Mediterranean environment?

Increasing air temperature due to changing climate is projected to decrease the length of the growing season, hasten vegetative development and maturation, and ultimately affect yield of many C3 crops. Previous multilocation trials highlighted strong relationships between thermal trends in the interval “beginning of flowering‐end of grain filling” and grain yield, and protein content in durum wheat (Triticum turgidum subsp. durum (Desf.) Husn.). With the aim to confirm these relationships, nine durum wheat genotypes, including old (Capeiti 8, Senatore Cappelli and Trinakria) and modern (Amedeo, Arcangelo, Mongibello, Simeto, and Svevo) varieties and a Sicilian landrace (Russello) were grown at three different sites representing a climate gradient in Sicily, Italy. Moreover, the effect of post‐anthesis heat stress on these durum wheats was further investigated in two contrasting environments: open‐field (control—C) and greenhouse heat stress (HS). HS shortened the interval “beginning‐end of flowering” of 1.5 days across genotypes, and the “end of flowering‐beginning of grain filling” and maturation of 4.9 days, with a range of 1 day in Arcangelo to 11 days in Cappelli. Advances in main phenophases significantly decreased kernel weight (KW) and grain yield (GY), whereas grain protein content (PC) increased. As expected, a negative relationship was observed between GY and PC, while positive relationships were found for GY and grain‐filling duration (GFD), and GY and KW. Genotypes responded differently to heat stress, as evidenced by the net photosynthesis, transpiration rate, instantaneous water use efficiency and dry matter accumulation in kernels. Genotypes were ranked according to the heat susceptibility index (HSI): Amedeo, Arcangelo, Capeiti 8, Svevo and Trinakria resulted heat‐tolerant. These varieties were characterized by an early trigger of dry matter accumulation in kernels under HS (Amedeo, Arcangelo and Trinakria), or showed similar length of the GFD (Capeiti 8) between environments. The multilocation trial confirmed a negative relationship between maximum temperatures and grain yield, and a positive relationship between minimum temperatures and protein content during grain–filling periods. Research focusing on agronomic strategies, phenology and breeding for tolerance to heat stress is of strategic importance to cope with the detrimental effect of global warming in semi‐arid climates.     

Remobilization of Nitrogen and Carbohydrate from Stems of Bread Wheat in Response to Heat Stress during Grain Filling

When wheat (Triticum aestivum L.) is grown under heat-stress conditions during grain filling, preanthesis stored total non-structural carbohydrates (TNC) and nitrogen (N) could serve as alternative source of assimilates. This study was performed to evaluate wheat genotypes for their ability to accumulate and remobilize TNC and N stored in their stem to support grain filling under heat stress. Eighteen genotypes were used for N remobilization study while nine of them were used for TNC remobilization study. They were grown in pots and placed in a vinyl house with the maximum temperature kept below 30 °C. Five days after anthesis (5DAA), half of the pots were taken to phytotrons where temperature was gradually increased and the maximum was set at 38 °C. Grain yield and grain weight decreased by about 35 % under heat stress. Significant differences were found among genotypes in percentage reduction in grain yield, grain weight, grain filling duration and harvest index because of heat stress. The N and TNC concentrations of the stem at 5DAA were significantly different among genotypes. Heat stress significantly reduced the N remobilization efficiency of most of genotypes. However, heat stress significantly increased TNC remobilization efficiency and significant variation were observed among genotypes. N remobilization efficiency across treatments significantly correlated with grain yield, grain weight, harvest index and grain filling duration. TNC at 5DAA negatively correlated with N at 5DAA and harvest index, but the TNC remobilization efficiency under heat stress positively correlated with mainstem grain yield, grain weight and harvest index. The rate of chlorophyll loss from flag leaf positively correlated with N and TNC remobilization efficiencies under heat stress suggesting a link between leaf senescence and remobilization efficiency. The results indicate that heat stress negatively affected grain yield, its components and N remobilization while it increased TNC remobilization because of the increasing demand for resources.     

Reduction in kernel weight as a potential indirect selection criterion for wheat grain yield under terminal heat stress

Heat is an important abiotic stress during wheat (Triticum aestivum L.) grain‐filling in South Asia. A study was undertaken to determine effectiveness of selection for reduction in 1000‐kernel weight (TKWR) under heat stress to increase grain yield. Selection was made for low and high TKWR and selected progenies were evaluated in timely and late seeded trials at two locations in Nepal in 2003. One thousand kernel weight (TKW), biomass yield, grain yield, harvest index (HI), grain‐filling duration (GFD) and area under spot blotch progress curve per day (AUDPC/day) were examined. The low and high TKWR groups did not differ significantly for TKW, biomass yield, grain yield, HI, days to heading, GFD and AUDPC/day under timely seeding. However, low TKWR lines showed higher TKW, biomass yield, grain yield, HI, and GFD and lower AUDPC/day than the lines with high TKWR under late seeding. Realized heritability estimates for TKWR ranged from 0.68 to 0.85. The findings show that selection for low TKWR could be used as an indirect selection criterion to identify high grain yielding lines under terminal heat stress.     

Mapping quantitative trait loci associated with grain filling duration and grain number under terminal heat stress in bread wheat (Triticum aestivum L.)

Terminal heat stress has the potential negative impact on wheat production across the world, especially in South Asia. Under the threat of terminal heat stress, wheat genotypes with stay green trait would suffer from high temperature stress during their long grain filling duration (GFD). The genotypes with short GFD would be advantageous. To identify quantitative trait loci (QTL) for heat tolerance, a RIL population of K 7903 (heat tolerant) and RAJ 4014 (heat sensitive) wheat genotypes was investigated under timely and late‐sown conditions. Heat susceptibility index of GFD, yield components and traits under late‐sown condition were used as phenotypic data for QTL identification. Stable QTLs associated with these traits were identified on chromosomes 1B, 2B, 3B, 5A and 6B. The LOD value ranged from 2.9 to 5.0 and the corresponding phenotyping variation explained ranged from 12.0–22%. QTL for heat susceptibility index for the grain filling duration were colocalized with QTL for productive tillers under late sown and GFD under late‐sown condition on chromosomes 1B and 5A, respectively. These genomic regions could be exploited for molecular wheat breeding programmes targeting heat tolerance.     

A Simple Drought‐Sensitive Model for Leaf : Stem Partitioning of Wheat

We investigated the leaf : stem partitioning of winter wheat (Triticum aestivum L. varieties ‘Dekan’ and ‘Batis’) with and without drought influence. Irrigated and drought‐stressed winter wheat, grown in a rainout shelter in 2009/10 and 2013/14, were sampled during shoot elongation phase at the experimental Farm Hohenschulen located in Northern Germany. The data set contains leaf (DM_L) and stem dry masses (DM_S), as well as measured water contents for several soil layers. A reduced relative dry matter allocation to leaves was observed under drought stress. Our results clearly show that, if simulated leaf : stem partitioning is not sensitive to drought, this will cause a positive bias in simulated leaf and a negative bias in simulated stem dry matter under water‐limited conditions. This is in accordance with previous studies which revealed that crop simulators often overestimate the impact of drought on light‐use efficiency, whereas the consequences on leaf area development are underestimated. However, the drought stress‐induced shift in leaf : stem partitioning is yet not considered by most common wheat crop simulators. Our aim was to fill the gap in simulation of drought stress implications on leaf area development. A simple allometric model for leaf : stem partitioning () was parameterized. Starting from the allometric leaf : stem relationship observed under optimum water supply, a correction term was introduced, which allows to adapt the partitioning to drought stress conditions. The lg‐transformed root‐weighted soil water potential in the rooting zone (lgψ_root, lg(hPa)), calculated as a function of measured water contents and simulated root distribution, was used as a drought stress indicator. The linear correction term assumes an increase of the stem fraction, proportional to the difference between lgψ_root and a drought stress threshold (pF_crit, lg(hPa)). The analysis revealed that the shift in allometric partitioning towards stem fraction starts with lgψ_root greater than 1.92 [lg(hPa)]. The slope of the relative increase of dry matter allocated to the stem fraction was determined with 0.26 [lg(hPa)^?1]. Both parameters of the correction term were found to be highly significant. Implications for crop modelling are discussed.     

Expression of seed storage proteins responsible for maintaining kernel traits and wheat flour quality in common wheat under heat stress conditions

Heat stress during grain filling has been documented to decrease wheat grain yield and quality in arid regions worldwide. We studied the effect of heat stress on wheat flour quality in heat tolerant cultivars to define the effects of heat stress on flour quality and to identify germplasm combining traits for heat tolerance and good flour quality. We studied the kernel phenotypic traits, the expression of seed storage proteins (SSPs), and the resulting flour quality under heat and normal conditions. Under heat stress, all cultivars yielded narrow-shaped seeds, and increased protein contents as compared to the control plants grown under normal conditions. The specific sedimentation values used to estimate the gluten quality varied between cultivars. We identified cultivars that could maintain good flour quality under heat stress conditions: ‘Imam’, which possessed the Glu-D1d allele responsible for the suitable bread-making; ‘Bohaine’, which displayed high expression level of SSPs; and ‘Condor’, which possessed slight variations in the ratio of each SSP under heat stress conditions. Combining the desirable traits from these cultivars could yield a wheat cultivar with heat tolerance and good flour quality.     

Molecular cytogenetic characterization and phenotypic evaluation of new wheat–rye lines derived from hexaploid triticale ‘Certa’ × common wheat hybrids

Hexaploid triticale contains valuable genes from both tetraploid wheat and rye and plays an important role in wheat breeding programmes. In order to explore the potential of hexaploid triticale ‘Certa’ in wheat improvement, two crosses were made using ‘Certa’ as female parent, and common wheat cultivars ‘Jinmai47’ (JM47) and ‘Xinong389’ (XN389) as male parents. The karyotyping of BCF_4:5 lines from Certa/JM47//JM47 and F_5:6 lines from Certa/XN389 was investigated using sequential fluorescence in situ hybridization (FISH). One 1B(1R) substitution line and five 1BL.1RS whole‐arm translocation lines were identified, one of which was found lacking ω‐secalin locus. Many structural alterations on wheat chromosomes were detected in the progeny. Great morphologic differences resulting from genetic variations were observed, among which the photosynthetic capability was increased while grain quality was slightly improved. Compared with both parents, the stripe rust resistance at adult stage was increased in lines derived from Certa/JM47//JM47, while it was decreased in lines derived from Certa/XN389. These newly developed lines might have the potential to be utilized in wheat improvement programmes.     

Drought Priming at Vegetative Growth Stage Enhances Nitrogen‐Use Efficiency Under Post‐Anthesis Drought and Heat Stress in Wheat

To study the effects of early drought priming at 5th‐leaf stage on grain yield and nitrogen‐use efficiency in wheat (Triticum aestivum L.) under post‐anthesis drought and heat stress, wheat plants were first exposed to moderate drought stress (drought priming; that is, the leaf water potential reached ca. ?0.9 MP a) at the 5th‐leaf stage for 11 days, and leaf water relations and gas exchange rates, grain yield and yield components, and agronomic nitrogen‐use efficiency (ANUE ) of the primed and non‐primed plants under post‐anthesis drought and heat stress were investigated. Compared with the non‐primed plants, the drought‐primed plants possessed higher leaf water potential and chlorophyll content, and consequently a higher photosynthetic rate during post‐anthesis drought and heat stress. Drought priming also resulted in higher grain yield and ANUE in wheat under post‐anthesis drought and heat stress. Drought priming at vegetative stage improves carbon assimilation and ANUE under post‐anthesis drought and heat stress and their combination in wheat, which might be used as a field management tool to enhance stress tolerance of wheat crops to multiple abiotic stresses in a future drier and warmer climate.     

Improving resistance against terminal drought in bread wheat by exogenous application of proline and gamma‐aminobutyric acid

Scarcity of water is a severe constraint, which hinders the wheat productivity worldwide. However, foliage application of osmoprotectants may be useful in reducing the drought‐induced yield losses in wheat (Triticum aestivum L.). In this study, potential of foliage applied osmoprotectants (proline, gamma‐aminobutyric acid) in improving the performance of bread wheat against terminal drought was evaluated. Both proline and gamma‐aminobutyric acid (GABA) were foliage applied at 50, 100 and 150 mg/L at anthesis stage (BBCH‐identification code‐ 61), in two bread wheat cultivars viz. Mairaj‐2008 and BARS‐2009. After 1 week of foliage application of these osmoprotectants, drought was imposed by maintaining the pots at 35% water holding capacity. Imposition of drought caused significant reduction in the grain yield of both tested bread wheat cultivars; nonetheless, foliage applied osmoprotectants at either concentration improved the chlorophyll contents, accumulation of proline, glycinebetaine and total soluble phenolics and reduced the malondialdehyde contents, which resulted in better stay green, maintenance of grain weight and grain number under drought stress, thus resulting in better grain yield, water‐use efficiency and transpiration efficiency in both wheat cultivars. However, foliage applied proline at 150 mg/L, and GABA at 100 mg/L was most effective than other concentrations of these osmoprotectants. Performance of cultivar Mairaj‐2008 was quite better than cultivar BARS‐2009. In crux, foliar application of proline and GABA at pre‐optimized rate can be opted as a shotgun approach to improve the performance of wheat under terminal drought.     

Leaf Photosynthesis During Grain Filling Under Mediterranean Environments: Are Barley or Traditional Wheat More Efficient Than Modern Wheats?

Barley is one of the most popular crops in dryland agricultural systems of Mediterranean areas, where it is assumed that barley, or traditional wheat cultivars, performs better than modern wheat under low‐yielding conditions. It was tested whether variations in net leaf photosynthetic rate (P_N) during grain filling provide any basis for the potential better performance of barley and traditional wheat compared to modern wheats in Mediterranean areas. Two groups of field experiments were conducted in Agramunt (NE Spain) during 2005/06 (06) and 2006/07 (07) growing seasons combining low and high nitrogen (N) availabilities under rain‐fed and irrigated conditions. Cultivars used in the first group of experiments were a traditional (Anza) and a modern (Soissons) wheat, whilst in a second group of experiments, a wheat (Soissons) and a barley (Sunrise) modern cultivars were used. Both wheat cultivars showed a similar P_N during grain filling but higher than that of the modern barley cultivar. Differences between species in P_N were maximized under high‐yielding conditions. There were no differences between cultivars in instantaneous water‐use efficiency. The barley cultivar showed a higher specific leaf area, but lower N content per unit of leaf area, than wheat. Photosynthetic nitrogen‐use efficiency was similar between the traditional and the modern cultivar but lower than barley. Decreases in P_N after anthesis were not exactly observable in SPAD measurements. In conclusion, we found no consistent differences between cultivars in terms of post‐anthesis photosynthetic activity to support the assumption of better performance under Mediterranean farm conditions of traditional wheat or barley against modern wheat.     

Application of natural plant extracts improves the tolerance against combined terminal heat and drought stresses in bread wheat

Drought and heat are among the main abiotic stresses causing severe damage to the cereal productivity when occur at reproductive stages. In this study, ten wheat cultivars were screened for combined heat and drought tolerance imposed at booting, heading, anthesis and post‐anthesis stages, and role of the foliage applied plant extracts was evaluated in improving the performance of differentially responding wheat cultivars under terminal heat and drought stresses. During both years, wheat crop was raised under ambient temperature and 70% water holding capacity (WHC) till leaf boot stage. The plant extracts (3% each) of sorghum, brassica, sunflower and moringa were foliage applied at booting, anthesis and post‐anthesis stage; and after one week of application of these plant extracts, combined heat and drought was imposed at each respective stage. Heat and drought stresses were imposed at each respective stage by placing pots in glass canopies with temperature of 4 ± 2°C above than the ambient temperature in combination with drought stress (35% WHC) until maturity. Combination of drought and heat stresses significantly reduced the performance of tested wheat cultivars; however, stress at the booting and heading stages was more damaging than the anthesis and post‐anthesis stages. Cultivars Mairaj‐2008 and Chakwal‐50 remained green with extended duration for grain filling, resulting in the maintenance of number of grains per spike and 100‐grain weight under stress conditions and thus had better grain yield and water‐use efficiency. However, in cultivars Fsd‐2008, and Shafaq‐2006, the combined imposition of drought and heat accelerated the grain filling rate with decrease in grain filling duration, grain weight and grain yield. Foliar application of all the plant extracts improved the wheat performance under terminal heat and drought stress; however, brassica extract was the most effective. This improvement in grain yield, water‐use efficiency and transpiration efficiency due to foliage applied plant extracts, under terminal heat and drought stress, was owing to better stay‐green character and accumulation of more soluble phenolics, which imparted stress tolerance as indicated by relatively stable grain weight and grain number. In crux, growing of stay‐green wheat cultivars with better grain filling and foliage application of plant extracts may help improving the performance of bread wheat under combined heat and drought stresses.     

Gas Exchange of Five Warm‐Season Grain Legumes and their Susceptibility to Heat Stress

Objective of this study was to compare the heat stress performance of four pulses from dry and hot areas (mungbeans, limabeans, and teparybeans and cowpeas) with that of soybeans. Two experiments were conducted in growth chambers, and data were pooled because results of both experiments were similar. Plants were raised up to flowering at 24/17 °C (day/night) and were then either exposed to these temperatures until maturity or stressed with 33/24 °C for 2 weeks starting at day 1 or 15 after onset of flowering (early vs. late stress). Before, during and after these stress intervals, gas exchange of representative upper leaves was examined; additionally, immediate effects of increasing leaf temperatures from 24 to 32 or 40 °C on chlorophyll fluorescence were assessed. Without heat stress rates of photosynthesis (P_n), and of transpiration (TR), stomatal and mesophyll conductance (g_s, g_m) and intrinsic transpiration efficiency (iTE) differed significantly among the five crops at each date. However, because of crop‐specific time‐courses ranking among unstressed crops was instable with time, so values were integrated or averaged over time. This procedure revealed high P_n potentials in mung‐ and teparybeans and high iTE values in limabeans compared to the other crops. Heat stress lowered P_n and g_s considerably, but increased TR in all five crops. Relative lowering of P_n during heat stress displayed a crop‐specific pattern with limabeans being least susceptible to both early and late heat stress, while cowpeas were highly susceptible to early stress. Effects on P_n were mainly attributable to lowering of g_s and only in part to g_m. The latter was supported by very small changes (<10 %) of various chlorophyll fluorescence signals shortly after raising leaf temperature to 32 °C in all species. However, in limabeans, a decreased electron transport rate (e‐rate, ?18 %) and an increased non‐photochemical quenching (Q_N, +16 %) pointed to an adaptive mechanism to avoid oxidative strains under heat. Leaf temperatures of 40 °C immediately provoked stronger changes in all fluorescence signals than 32 °C; substantial damages at 40 °C were indicated by effective quantum yield, photochemical quenching and ratio of fluorescence decrease in mungbeans and low ones in cowpeas and soybeans. Nevertheless, some adaptive responses of e‐rates and Q_N were observed in all crops and were most expressed in limabeans.     

Effect of two-week heat stress during grain filling on stem reserves,senescence, and grain yield of European winter wheat cultivars

To examine the extent to which heat stress during grain filling impacts on the development and yield of winter wheat (Triticum aestivum L.), a 3-year field experiment was conducted on a loess soil with high water holding capacity in the North German Plain. Thirty-two mostly European winter wheat cultivars were exposed to heat stress in a mobile foil tunnel with maximum air temperatures of 45.7, 45.4, and 47.2°C in 2015, 2016, and 2017, respectively. The 14-day post-anthesis heat stress treatment caused an average 57.3% grain yield reduction compared to a close-by non-stressed control. The proportion of green crop area after the heat stress phase varied from 7% to 98% in 2016 and from 37% to 94% in 2017. The green crop area percentage did not significantly correlate with grain yield, indicating that the delayed senescence of stay-green phenotypes offers no yield advantage under terminal heat stress. The water soluble carbohydrate (WSC) concentration of the stems at crop maturity varied between 6 and 92 g/kg dry matter, showing that the genotypes differed in their efficiency at using the stem carbohydrate reserves for grain filling under heat stress. The stem WSC concentration correlated positively with the beginning of anthesis (r = 0.704; p < .001) but negatively with the grain yield (r = −0.431; p < .05). For heat tolerance breeding, the stem reserve strategy, i. e. the rapid and full exhaustion of the temporary carbohydrate storage therefore seems more promising than the stay-green strategy.